Long-term persistence of aspen – a key host for many threatened species – is endangered in old-growth conservation areas in Finland

Large, dead and dying European aspens (Populus tremula L.) host many threatened species in Fennoscandian boreal forests. Large aspen trees have mostly disappeared and are being harvested from the managed forests that cover 95% of the forest area in Finland. Due to the small area protected (4.1%), the aspen-associated species may encounter major difficulties in the protected areas if aspen trees disappear due to natural forest succession. The availability of aspens was assessed in the old-growth conservation area network in eastern Finland. We mapped all the living and dead aspens in 15 protected old-growth forests. The total number of counted trees was 32 903 individuals. Current amounts of living (2.7 m³/ha) and especially dead aspens (2.8 m³/ha) in the protected areas were higher than in the surrounding managed forests (1.1 and 0.1 m³/ha for living and dead trees, respectively). However, while saplings (dbh<5 cm) occur in most of the areas (12 individuals/ha on average) they survive poorly and young aspen cohorts (5 cm <dbh<15 cm) are lacking or are very rare. The most likely reason for the poor sapling survival is high browsing pressure by the mammalian herbivores, especially the moose. The moose population has increased many times in Finland during the past decades. The poor regeneration of aspens implies that the value of the old-growth conservation areas for aspen-associated species will face a serious bottleneck within a few decades when the currently middle-aged tree cohorts disappear. If the current high browsing pressure and lack of natural disturbances continue the obligatory aspen-associated species may disappear both locally and regionally from the network of the protected areas.     

Clonal dynamics in western North American aspen (Populus tremuloides)

Clonality is a common phenomenon in plants, allowing genets to persist asexually for much longer periods of time than ramets. The relative frequency of sexual vs. asexual reproduction determines long‐term dominance and persistence of clonal plants at the landscape scale. One of the most familiar and valued clonal plants in North America is aspen (Populus tremuloides). Previous researchers have suggested that aspen in xeric landscapes of the intermountain west represent genets of great chronological age, maintained via clonal expansion in the near absence of sexual reproduction. We synthesized microsatellite data from 1371 ramets in two large sampling grids in Utah. We found a surprisingly large number of distinct genets, some covering large spatial areas, but most represented by only one to a few individual ramets at a sampling scale of 50 m. In general, multi‐ramet genets were spatially cohesive, although some genets appear to be fragmented remnants of much larger clones. We conclude that recent sexual reproduction in these landscapes is a stronger contributor to standing genetic variation at the population level than the accumulation of somatic mutations, and that even some of the spatially large clones may not be as ancient as previously supposed. Further, a striking majority of the largest genets in both study areas had three alleles at one or more loci, suggesting triploidy or aneuploidy. These genets tended to be spatially clustered but not closely related. Together, these findings substantially advance our understanding of clonal dynamics in western North American aspen, and set the stage for a broad range of future studies.     

Molecular and dendrochronological analysis of natural root grafting in Populus tremuloides (Salicaceae)

Trembling aspen (Populus tremuloides) is a clonal tree species, which regenerates mostly through root suckering. In spite of vegetative propagation, aspen maintains high levels of clonal diversity. We hypothesized that the maintenance of clonal diversity in this species can be facilitated by integrating different clones through natural root grafts into aspen's communal root system. To verify this hypothesis, we analyzed root systems of three pure aspen stands where clones had been delineated with the help of molecular markers. Grafting between roots was frequent regardless of their genotypes. Root system excavations revealed that many roots were still living below trees that had been dead for several years. Some of these roots had no root connections other than grafts to living ramets of different clones. The uncovered root systems did not include any unique genotypes that would not occur among stems. Nevertheless, acquiring roots of dead trees helps to maintain extensive root systems, which increases the chances of clone survival. Substantial interconnectivity within clones as well as between clones via interclonal grafts results in formation of large genetically diverse physiological units. Such a clonal structure can significantly affect interpretations of diverse ecophysiological processes in forests of trembling aspen.     

Clonal and spatial genetic structures of aspen (Populus tremuloides Michx.)

To portray aspen clonal and spatial genetic structures, we mapped and genotyped trees in two 1-ha plots, each containing three aspen cohorts originating from fire or subsequent secondary disturbances. We used four microsatellite loci to identify aspen clones and increment core analysis to determine tree age. Clonal dimensions were measured by the maximum distance between two ramets and the number of ramets per genet. Standard normal deviate (SND) was used to assess the spatial distribution of aspen genets and cohorts, and multivariate spatial genetic autocorrelations to assess the spatial distribution of aspen genetic variation. Most aspen genets consisted of only one ramet (> 75%). Median clonal dimensions were 19 and 29 m (maxima: 104 and 72 m in the two plots). No segregation was observed between clones. Aspen cohorts were spatially segregated but trees were spatially aggregated within old and medium-aged cohorts. In contrast, trees were more randomly distributed within the youngest cohorts. This coincided with a spatial genetic autocorrelation at small scales (up to 30 m) in the older cohorts and a more random genetic distribution in the youngest ones. Our results suggest that aspen spatial genetic structuring reflects the spatial patterns produced by the regeneration of discrete cohorts at different stages of succession. Vegetative reproduction leads to aspen genetic spatial structuring at small scales (few metres) until midsuccession. However, as the stand gets older, the spatial distribution of aspen trees and genetic structure evolve from a structured pattern to a more random one under a gap disturbances regime.     

Spatial Variation of Bird Species in Landscapes Dominated by Old-growth Forests in Northern Boreal Finland

An important issue in conservation planning is to study the distribution and abundance patterns of species in natural landscapes. Information of concordant and clumped species distribution pattertns can enhance attempts to focus conservation efforts in sites/areas preferred by various species. We compared bird species abundance, community composition and species distribution patterns in three large old-growth forest areas (size 40–120 km²) in northern boreal Finland on the basis of quantitative bird censuses. Total bird density and species composition, and mean density and variances of the most abundant species were highly similar between the three areas. Most of the bird species were distributed randomly within the areas and species preferring old-growth forests showed compensatory density variation without preference for any sub-area. Mean density of species preferring old-growth forests was significantly higher in the studied large old-growth forests than in smaller old-growth forests and was about threefold higher than regional density of species in a predominantly managed landscape matrix. Bird species preferring old-growth forests are probably not dependent on a certain specific part of a virgin boreal forest landscape but rather on the overall size of the high-quality and diverse old-growth forest area.     

Fallen retention aspen trees on clear-cuts can be important habitats for red-listed polypores: a case study in Finland

Green-tree retention is a relatively new forestry application, which aims at decreasing the negative effects of clear-cut logging on forest biodiversity. In this study, the value of retained aspens in maintaining diverse assemblages of wood-decaying fungi (polypores; Basidiomycota) on clear-cuts was investigated, after the retention trees had died, fallen and started to decay. A total of 110 fallen aspen trunks were investigated on clear-cuts and within old-growth forests in eastern Finland, southern boreal zone; and 499 records of polypores belonging to 46 species were made. The intermediately decayed trunks on a clear-cut area hosted more species and more red-listed species than did trunks within forests. Most of the polypore species with more than two records were found in both habitats. These results suggest that many aspen-associated polypores are able to live and reproduce in sun-exposed habitats, if the quality and quantity of dead wood fulfill the species-specific requirements. This unexpected result, however, may be partly due to the exceptionally great abundance of aspen in the study area. Furthermore, in the long term, the local benefits of fallen retention trees can be limited, unless the local continuity of large aspens, both living and dead, is ensured.     

Consequences of forest fragmentation for polyporous fungi at two spatial scales

Greatly reduced area of old-growth forests and the very low amount of dead wood in managed forests in northern Europe have caused a marked decline in the populations of saproxylic species. It is less clear at which spatial and temporal scales these adverse changes are taking place, and more information is needed to reliably predict which species are especially sensitive to loss and fragmentation of habitat. Here we compare species richness, incidence of occurrence in forest fragments, and abundance of polyporous fungal species and species groups between two regions in Finland with contrasting histories of forestry and a marked difference in the amount and spatial configuration of old-growth forests. We also analyse the consequences of increasing loss of connectivity on the presence and abundance of polypores in a study region with a documented short-term history of old-growth fragmentation. Our results show that the species number, incidence of occurrence, and abundance of especially the rare, threatened, and near-threatened species are much lower in the old-growth fragments in Häme in southern Finland in comparison with Kuhmo in eastern Finland, most probably because of the longer history of intensive forestry in Häme. Among the rare species, the species that show the greatest difference between the two regions (at the scale of 500 km) also tended to respond most strongly to the more recent forest fragmentation within the study region in Kuhmo (at the scale of 50 km). Polypores associated with spruce seem to be more strongly affected by forestry than species associated with pine, possibly reflecting the differences in the natural dynamics of spruce-dominated and pine-dominated forests.     

Selectivity by moose vs the spatial distribution of aspen: a natural experiment

Patch use theory predicts that herbivores perceive food as patches and spend more time in high quality patches, i.e. feeding sites providing a high net rate of intake of energy and/or limiting nutrients. The herbivores should accordingly not discriminate among food items in such high quality patches, and food items should thus be eaten in proportion to availability. In contrast, classical diet theory assumes food selection to take place at the level of individual plants, and predicts that the forager should concentrate on the most profitable item until availability drops below some critical threshold.
Here we address how the spatial distribution of European aspen Populus tremula , a highly preferred browse species, affects the selectivity by moose Alces alces at the patch and the tree level. The study was performed in a managed boreal forest landscape in coastal northern Sweden, where aspen occurs highly aggregated almost exclusively in discrete patches. We compared moose' selectivity for aspen and browsing intensity on aspen ramets and other browse species in aspen patches versus at randomly located sites.
Random sites and aspen stands were utilised equally by moose in terms of overall use of forage. There was no difference in total coverage of forage species and relative moose density. Selectivity for aspen was stronger at random sites than at aspen sites, and the browsing intensity on aspen was similar. We conclude that moose did not perceive aspen stands as discrete patches, and used aspen ramets more in accordance with diet theory. These findings agree with the idea that large generalist herbivores strive to maintain a mixed and balanced diet, causing rare species to be over-utilised (negative frequency-dependent food selection). By such selective feeding, moose may reinforce the aggregated distribution of aspen in the managed boreal forest landscape.     

Leaf litter and the small-scale distribution of carabid beetles (Coleoptera, Carabidae) in the boreal forest

Management practices favoring conifers at the expense of deciduous tree species, and the eradication of deciduous trees, especially aspen Populus tremula , from managed forests have resulted in population declines in several species in Fennoscandia. In addition to species depending on decaying wood of deciduous trees, earlier evidence suggests that leaf litter, especially that of aspen, is favored by many carabid species. We ran a four-year experiment in order to compare carabid assemblages of unchanged forest floor with artificially created leaf-litter plots in central Finland. A total of 18 plots (5 m in diameter) were established in three forest stands without aspen a few kilometers apart. Each stand had 3 litter plots (litter added) and 3 control plots, Pre-treatment samples were compared with those collected alter litter addition.
The litter addition affected the carabid-assemblage structure by increasing the catches of some species and decreasing the catch of one species. The number of carabid species was similar in control and litter plots. The litter effect was smaller than variation among forest stands and year-lo-year fluctuations. There was a strong temporal constancy among the plots: 'rich' plots remained 'rich' from year to year and similarly, 'poor' plots remained 'poor'.
The significant influence of leaf litter on carabid abundance can be attributable to both abiotic factors (microenvironmental conditions, especially humidity and temperature), and biotic ones (changes in niche structure, improved food supply). Leal litter seems to have an effect on carabid distribution patterns, and deciduous trees scattered among conifers are likely to be of importance on carabid fauna in boreal forests.     

Spatial pattern in the occurrence of the lichen Lobaria pulmonaria in managed and virgin boreal forests

Two populations of the lichen Lobaria pulmonaria. growing on aspens and goat willows in 12 and 20 km² study areas of boreal forest in Finland, were surveyed thoroughly to investigate the factors influencing the spatial distribution of the lichen. In one study area, where forestry has been intensive and old-growth forest is highly fragmented, L, pulmonaria was sparse and grew mostly on willows. In contrast, a large and continuous virgin forest area supported a higher incidence of L. pulmonaria. with the lichen being common on both aspens and willows. In both study areas, the distributions of aspen and willow were clumped over the scales of 100-1000 m. The spatial pattern of L pulmonaria was more clumped in the managed forest than in the virgin forest. The reduced incidence of the lichen on aspens in the managed area was attributed to a disruption of habitat continuity and small average tree size. There was no comparable reduction in the incidence on willows, probably because the willow had a very aggregated distribution in the managed area, which probably facililated local colonization of the lichen. Presence of the lichen was significantly related to size-corrected local density of aspen and willow trees as well as to spatial connectivity to neighboring lichen-occupied trees.     

Comparison of phenotypic and genetic clone delineation in quaking aspen, <Emphasis Type="Italic">Populus tremuloides</Emphasis>

Key message

Clonal delineation at nuclear microsatellites and phenotypic traits showed high correspondence and revealed an important role of both sexual and clonal reproduction for stand genetic structure.

Abstract

Quaking aspen (Populus tremuloides Michx.) grows throughout the northern and central portions of North America. Reproduction occurs both sexually via seeds and clonally from root suckers. Clonal delineation using morphological/phenological traits, and more recently, highly variable nuclear microsatellites have shown considerable variation in the size of clonal assemblies, and the relative importance of sexual versus clonal reproduction across the species range. In order to provide reliable estimates of genet size (N/G; ramets per sampled genet) and genotypic diversity (G/N; genets/ramets), and to compare genetic and phenotypic clone delineation, we characterized 181 sampled stems (ramets) at seven nuclear microsatellites, and morphological and phenological traits from six clones (genet size ≥11). Genotypic diversity was moderate (G/N = 0.18) and within the range reported in other studies across North America. Multivariate statistics revealed a high correspondence between genetic and phenotypic clone delineation, both with and without predefined genetic groups (94.2 %, 81.7 %). Moderate average genet size (5.6 ramets per genet) and the occurrence of genetically distinct single-ramet genets surrounded by larger genets suggested intermediate levels of sexual reproduction contributing to the genetic structure of this stand. Significant differences among genets were found for phenological and morphological traits such as bark thickness and leaf shape. However, most clones showed no significant differences in diameter growth which was likely caused by poor drainage in this high clay soil that inhibited the expression of genetic differences in growth.

     

Loss of whole-tree hydraulic conductance during severe drought and multi-year forest die-off

Understanding the pathways through which drought stress kills woody vegetation can improve projections of the impacts of climate change on ecosystems and carbon-cycle feedbacks. Continuous in situ measurements of whole trees during drought and as trees die hold promise to illuminate physiological pathways but are relatively rare. We monitored leaf characteristics, water use efficiency, water potentials, branch hydraulic conductivity, soil moisture, meteorological variables, and sap flux on mature healthy and sudden aspen decline-affected (SAD) trembling aspen (Populus tremuloides) ramets over two growing seasons, including a severe summer drought. We calculated daily estimates of whole-ramet hydraulic conductance and modeled whole-ramet assimilation. Healthy ramets experienced rapid declines of whole-ramet conductance during the severe drought, providing an analog for what likely occurred during the previous drought that induced SAD. Even in wetter periods, SAD-affected ramets exhibited fivefold lower whole-ramet hydraulic conductance and sevenfold lower assimilation than counterpart healthy ramets, mediated by changes in leaf area, water use efficiency, and embolism. Extant differences between healthy and SAD ramets reveal that ongoing multi-year forest die-off is primarily driven by loss of whole-ramet hydraulic capability, which in turn limits assimilation capacity. Branch-level measurements largely captured whole-plant hydraulic limitations during drought and mortality, but whole-plant measurements revealed a potential role of other losses in the hydraulic continuum. Our results highlight the importance of a whole-tree perspective in assessing physiological pathways to tree mortality and indicate that the effects of mortality on these forests’ assimilation and productivity are larger than expected based on canopy leaf area differences.     

Habitat associations of <Emphasis Type="Italic">Agathidium pulchellum</Emphasis>, an endangered old-growth forest beetle species living on slime moulds

The beetle genus Agathidium is the largest insect group documented that principally feeds on slime moulds. Agathidium pulchellum, one of the rarest Agathidium species in Europe, is listed in the EU’s Habitats Directive. We studied the habitat associations of A. pulchellum in 44 sites located in old-growth and managed forests in eastern Finland. Agathidium pulchellum occurred exclusively on the slime-mould species Trichia decipiens. The host was associated with mid-decayed aspen, spruce and birch logs, and its incidence grew with both increasing log diameter and stand-level log density of spruce and aspen. We also observed that even if its host was present, the beetle was absent from sites with less than 80 aspen and spruce logs per hectare. All sites with A. pulchellum were natural forests of high conservation value. Our results show that it is possible to systematically survey the occurrence of A. pulchellum.     

Infestation and hydraulic consequences of induced carbon starvation

Drought impacts on forests, including widespread die-off, are likely to increase with future climate change, although the physiological responses of trees to lethal drought are poorly understood. In particular, in situ examinations of carbon starvation and its interactions with and effects on infestation and hydraulic vulnerability are largely lacking. In this study, we conducted a controlled, in situ, repeated defoliation experiment to induce carbon stress in isolated trembling aspen (Populus tremuloides) ramets. We monitored leaf morphology, leaves per branch, and multitissue carbohydrate concentrations during canopy defoliation. We examined the subsequent effects of defoliation and defoliation-induced carbon stress on vulnerability to insect/fungus infestation and hydraulic vulnerability the following year. Defoliated ramets flushed multiple canopies, which coincided with moderate drawdown of nonstructural carbohydrate reserves. Infestation frequency greatly increased and hydraulic conductivity decreased 1 year after defoliation. Despite incomplete carbohydrate drawdown from defoliation and relatively rapid carbohydrate recovery, suggesting considerable carbohydrate reserves in aspen, defoliation-induced carbon stress held significant consequences for vulnerability to mortality agents and hydraulic performance. Our results indicate that multiyear consequences of drought via feedbacks are likely important for understanding forests' responses to drought and climate change over the coming decades.     

Importance of habitat patch size for occupancy and density of aspen-associated saproxylic beetles

In Fennoscandian boreal forests, aspen (Populus tremula) is one of the most important tree species for biodiversity. In this study we explore how occupancy and density of beetles associated with dead aspen are related to habitat patch size and connectedness in a 45,000 ha boreal managed forest landscape in central Sweden. Patch size was estimated as amount of breeding substrate and connectedness as crown cover of living aspen in the surrounding landscape. The beetles were sampled by sieving of bark or by inspection of species-characteristic galleries in 56 patches with dead aspen. Six of nine aspen-associated species (Xylotrechus rusticus, Ptilinus fuscus, Mycetophagus fulvicollis, Cyphaea curtula, Homalota plana and Endomychus coccineus) showed a positive significant relationship between habitat patch size and occupancy. For all these species, except C. curtula, there was also a significant positive relationship between patch size and density. Connectedness was not retained as a significant variable in the analyses. Species not defined as aspen-associated constituted a significantly larger proportion of the total density of individuals of saproxylic beetles in smaller habitat patches than in larger patches. Richness of aspen-associated species was positively related to habitat patch size. Efforts in the managed forest should be directed towards preserving and creating larger patches of living and dead aspen trees and increasing the amount of aspen at the landscape level.     

Dispersal ecology of the endangered woodland lichen Lobaria pulmonaria in managed hemiboreal forest landscape

Changes in the forest management practices have strongly influenced the distribution of species inhabiting old-growth forests. The epiphytic woodland lichen Lobaria pulmonaria is frequently used as a model species to study the factors affecting the population biology of lichens. We sampled 252 L. pulmonaria individuals from 12 populations representing three woodland types differing in their ecological continuity and management intensity in Estonia. We used eight mycobiont-specific microsatellite loci to quantify genetic diversity among the populations. We calculated the Sørensen distance to estimate genetic dissimilarity among individuals within populations. We revealed that L. pulmonaria populations have significantly higher genetic diversity in old-growth forests than in managed forests and wooded meadows. We detected a significant woodland-type-specific pattern of genetic dissimilarity among neighbouring L. pulmonaria individuals, which suggests that in wooded meadows and managed forests dominating is vegetative reproduction. The vegetative dispersal distance between the host trees of L. pulmonaria was found to be only 15–30 m. Genetic dissimilarity among individuals was also dependent on tree species and trunk diameter. Lobaria pulmonaria populations in managed forests included less juveniles compared to old-growth forests and wooded meadows, indicating that forest management influences life stage structure within populations. We conclude that as intensive stand management reduces the genetic diversity of threatened species in woodland habitats, particular attention should be paid to the preservation of remnant populations in old-growth habitats. Within managed habitats, conservation management should target on maintenance of the stand’s structural diversity and availability of potential host trees.     

Aspen (<Emphasis Type="Italic">Populus tremuloides</Emphasis>) stands and their contribution to plant diversity in a semiarid coniferous landscape

We conducted a field study to determine the relative contributions of aspen (Populus tremuloides), meadow, and conifer communities to local and landscape-level plant species diversity in the Sierra Nevada and southern Cascade Range, northeastern California, USA. We surveyed plant assemblages at 30 sites that included adjacent aspen, conifer, and meadow communities across a 10,000-km² region. We statistically investigated patterns in local and landscape-scale plant diversity within and among the three vegetation types. Summing across sites, aspen stands supported more plant species overall and more unique plant species than either meadow or conifer communities. Local richness and diversity did not differ between aspen and meadow plots; conifer forest plots were significantly lower in both measures. Heterogeneity in species composition was higher for aspen forest than for meadows or conifer forest, both within sites and between sites. Plant communities in aspen stands shared less than 25% of their species with adjacent vegetation in conifer and meadow plots. Within aspen forest, we found a negative relationship between total canopy cover and plant diversity. Our results strongly support the idea that plant communities of aspen stands are compositionally distinct from adjacent meadows and conifer forest, and that aspen forests are a major contributor to plant species diversity in the study region. Current patterns of aspen stand succession to conifer forest on many sites in the semiarid western US are likely to reduce local and landscape-level plant species diversity, and may also have negative effects on other ecosystem functions and services provided by aspen forest.     

Microsatellite analysis of clonality and individual heterozygosity in natural populations of aspen <Emphasis Type="Italic">Populus tremula</Emphasis> L.: Identification of highly heterozygous clone

Aspen Populus tremula L. (Salicaceae) is the fast-growing tree species of environmental and economic value. Aspen is capable of reproduction by both seeds and vegetative means, forming root sprouts. In an adult stand, identification of ramets of one clone among the trees of seed origin based on their morphology is difficult. A panel of 14 microsatellite loci developed for individual identification of aspen was applied for the clonal structure analysis in four natural aspen stands of the European part of Russia: Moscow and Voronezh oblasts, the Mari-El Republic, and the Republic of Tatarstan. In 52 trees from the Moscow sample, 41 multilocus genotypes were identified; in the Voronezh sample, among 30 individuals, 25 different genotypes were detected; and in the sample from Mari-El, 32 trees were represented by 13 genotypes. In the stand from Sabinsky Forestry, Tatarstan, all of the examined 29 trees were represented by a single genotype. The ancestral tree carrier of this genotype which was the most heterozygous (0.929) among all studied aspen individuals (sample mean, 0.598) obviously has spread over a large territory during several cutting and reproduction cycles, currently occupying the area of 2.2 ha. For aspen, usually suffering from Aspen trunk rot, such high viability is evidence of resistance to the main pathogens. The revealed superclone deserves further study with karyological methods and flow cytometry to determine ploidy level and analysis of the growth rate and the quality of wood for possible use in plantation forest production.     

Genetic mosaics of ecosystem functioning across aspen-dominated landscapes

Genetic diversity is the foundation of all biodiversity, and the genetic variation within species is increasingly recognized as being important to ecosystem level processes. Recent research demonstrates that plant genotype influences above- and belowground communities as well as basic ecosystem functions. However, the extent to which plant genotypes create spatial mosaics of genetically mediated ecosystem processes in natural forests is uncertain. We use Populus tremuloides as a model system to demonstrate the importance of plant genotype on carbon and nitrogen cycling in natural systems. We identified 24 distinct P. tremuloides clones with multiple ramets across 25 km² in southern Wisconsin, United States, using microsatellite makers. We then sampled clone leaf chemistry and belowground nutrient content and microbial extracellular enzyme activity. Aspen-induced variation in belowground carbon and nitrogen content, and microbial activity, varied widely among clones. Variation in green leaf chemistry and belowground microbial activity were correlated with genetic distance among clones, such that more genetically distant clones created more divergent patches of ecosystem processes. These data suggest that aspen genotypes create spatial mosaics of genetically mediated ecosystem functioning across natural landscapes and can therefore have evolutionary consequences for co-occurring species.     

Nitrogen translocation between clonal mother and daughter trees at a grassland–forest boundary

There is abundant evidence from short-term experiments using herbs that nutrients can be translocated from mother ramets to daughter ramets, but there is little long-term evidence from woody plants. Here, we examine translocation in field populations of a clonal tree over two growing seasons. We applied ¹⁵N to mothers or daughters in clones of Populus tremuloides at the northern edge of the North American Great Plains, where mother ramets form closed-canopy stands on relatively nutrient-rich soils, and daughter ramets occur nearby in relatively nutrient-poor grasslands. Unlabeled daughters in clones with labeled mothers had δ¹⁵N values significantly greater than those in unlabeled clones, confirming translocation from mothers to daughters. However, unlabeled mothers in clones with labeled daughters also had δ¹⁵N values significantly greater than those in unlabeled clones, indicating translocation from daughters to mothers. Further, the total foliage accumulation of added ¹⁵N was significantly (c. 10×) greater in mothers than in daughters, suggesting that more N was translocated from daughters to mothers, than from mothers to daughters. Thus, ¹⁵N moved both from mothers to daughters and from daughters to mothers, with net flow toward mothers. Because long-lived woody ramets in the field face nutrient competition from other ramets, interspecific neighbors, and soil microbes, the environmental availability of nutrients for uptake may be low for both mother and daughter ramets, causing translocation within a clone to be toward larger ramets with greater demand.